NORTH-WESTERN JOURNAL OF ZOOLOGY 8 (2): 205-214 ©NwjZ, Oradea, Romania, 2012 Article No.: 121114 http://biozoojournals.3x.ro/nwjz/index.html

Description of all developmental stages of Allochernes peregrinus Lohmander, 1939 (Pseudoscorpiones, )

Jana CHRISTOPHORYOVÁ1*, Katarína KRAJČOVIČOVÁ1, Miroslav KRUMPÁL1 and Karel TAJOVSKÝ2

1. Department of Zoology, Faculty of Natural Sciences, Comenius University, Mlynská dolina B–1, SK–842 15 Bratislava, Slovakia. 2. Biology Centre, Academy of Sciences of the Czech Republic, Institute of Soil Biology, Na Sádkách 7, CZ–370 05 České Budějovice, Czech Republic. * Corresponding author, e-mail: [email protected]

Received: 11. November 2011 / Accepted: 8. April 2012 / Available online: 26. May 2012 / Printed: December 2012

Abstract. An illustrated description of all developmental stages of the species Allochernes peregrinus Lohmander, 1939 is provided based on morphology and morphometrics. During the research, 48 specimens of Allochernes peregrinus were studied, with 46 of them collected in the Czech Republic and two in Slovakia. The analysis of female materials updated the known variability of morphological and morphometric characters, and the first descriptions of males and nymphs are given here. The main determination characters of the species are compared with those of other species in the genus Allochernes occurring mainly in neighbouring European countries. Habitat distribution of A. peregrinus is briefly discussed.

Keywords: Pseudoscorpiones, Allochernes peregrinus, Czech Republic, Slovakia, morphology, morphometric analysis, taxonomy.

Introduction (1972) noted that a single specimen was not suffi- cient for indisputable assignment to this species. represent an order of a Some adults were later recorded in Central Europe medium diversity. Europe represents the best without any notes concerning their determination documented continent, with a known pseudo- characters – in Germany (Drogla 1983, Platen et al. scorpion fauna of about 760 species (Harvey 2007). 1995, Drogla & Lippold 2004), Slovakia (Krum- The genus Allochernes Beier, 1932 includes 31 spe- pálová & Krumpál 1993, Christophoryová & cies and 2 subspecies, and it is spread in North Af- Krumpál 2007, 2010), Czech Republic (Ducháč rica and widespread in Asia and Europe (Harvey 1995, 1999) and Austria (Mahnert 2004). Several 2009). The pseudoscorpion Allochernes peregrinus morphological and morphometric features of Lohmander, 1939 is distributed across several adults and tritonymphs were presented in the un- countries of Central and Northern Europe (Fig. published theses of DeVore-Scribante (1999) and 1A) – Austria, Czech Republic, Germany, Hun- Ducháč (2002). Two females of Allochernes aff. gary, Poland, Slovakia, Switzerland and Sweden, peregrinus were collected in Azerbaijan by Dash- and in the USA (DeVore-Scribante 1999, Harvey damirov (1999). Until present only isolated records 2009). The species was described by Lohmander of A. peregrinus were known from the Czech Re- (1939) from Sweden based on one phoretic female public and Slovakia (Krumpálová & Krumpál attached to the leg of the fly Delia floralis (Fallén, 1993, Ducháč 1995, 2002, Christophoryová & 1824) (syn. Hylemyia floralis). Almost 30 years later, Krumpál 2007, 2010). Therefore this species was Rafalski (1967) recorded specimens of this species listed in the Red Data Book of the Czech Republic in Poland in deeper soil layers and under the as vulnerable (Šťáhlavský & Ducháč 2005). stones sunken into soil, but he did not specify Incomplete (and unpublished) description of which developmental stages were found. Mahnert males of A. peregrinus, together with the descrip- (1983) gave more data on morphological and mor- tion of females based only on a few specimens phometric characteristics of one female specimen poorly reflecting intraspecific variability and miss- sampled in Hungary. The first male of A. peregri- ing description of nymphs, led us to elaborate nus was found by Muchmore (1972, 1990) in beech these materials in more detail including i) descrip- leaf litter in New Hampshire, USA. Muchmore tion of all developmental stages of A. peregrinus in 206 J. Christophoryová et al. detail, ii) variability of morphological and mor- sented seven forest plant associations characteristic for phometric characteristics, and iii) summarization the given area: thermophilous oak (TO), mesophilous of all available data concerning distribution and oak-hornbeam (MOH), thermophilous oak-hornbeam (TOH), acidophilous oak (AO), basiphilous oak (BO), habitat preferences in Europe. plantations of deciduous (DP) and coniferous (CP) trees.

2 At each site, five soil samples (area of each 625 cm , depth Materials and Methods ca 10 cm) were taken in spring and autumn and speci- mens were heat extracted from samples in modified In total 48 specimens of Allochernes peregrinus were exam- Kempson apparatus. Five pitfall traps (each with diame- ined (15 females, 15 males, 10 tritonymphs, 6 deuto- ter 9 cm and formaldehyde killing-preserving solution) nymphs, 2 protonymphs). Forty-six of them were sam- with a year continual exposition and monthly sampling pled in the Czech Republic in fragmented forest land- intervals were used at each study site. Two adults from scape of the Český kras Protected Landscape Area in Cen- Slovakia (Fig. 1B) were included to the morphological tral Bohemia (Fig. 1B). During 2008–2009 soil sampling and morphometric analysis, findings of which were al- and pitfall trapping were used as methods in a set of se- ready published (Christophoryová & Krumpál 2007, lected forest fragments. Individual sampling sites repre- 2010).

A

B

Figure 1. Distribution of Allochernes peregrinus in Europe (A) with the detail of distribution in the Czech Republic and Slovakia (B). Square indicates the type locality (Lohmander 1939), triangles indicate known and published records (Mahnert in litt., Rafalski 1967, Drogla 1983, Mahnert 1983, Krumpálová & Krumpál 1993, Ducháč 1995, Drogla & Lippold 2004, Christophoryová & Krumpál 2007, 2010) and circles indicate new records of species distribution. Description of developmental stages of Allochernes peregrinus 207

The faunistic data: Czech Republic, Český Kras, lo- row distinct, posterior furrow indistinct and situ- cality Strážište (leg. K. Tajovský, A. Tenčík) – formalde- ated close to posterior margin of carapace. Cheli- hyde traps situated in: MOH (49°92'N, 14°12'E, 422 m cerae (Fig. 2B): small, slightly sclerotized, 5 setae a.s.l.), 30.4.– 28.5.2008, 1 female; BO (49°92'N, 14°11'E, 432 on cheliceral hand, one on cheliceral movable fin- m a.s.l.), 30.4.–28.5.2008, 2 females; BO (49°92'N, 14°11'E, 432 m a.s.l.), 28.5.–25.6.2008, 1 female; soil samples from: ger; movable cheliceral finger with slender well- DP (49°91'N, 14°09'E, 234 m a.s.l.), 5.5.2009, 2 deuto- developed galea, main stalk with approximately nymphs; DP (49°91'N, 14°12'E, 352 m a.s.l.), 5.5.2009, 1 5–6 short terminal rami; cheliceral rallum of 3 deutonymph; MOH (49°91'N, 14°12'E, 415 m a.s.l.), blades, first distal blade of cheliceral rallum den- 5.5.2009, 5 males; MOH (49°92'N, 14°12'E, 422 m a.s.l.), ticulate anteriorly; small, largely unsclerotized 5.5.2009, 2 males, 5 tritonymphs; TOH (49°92'N, 14°12'E, teeth situated on both movable and fixed finger of 405 m a.s.l.), 5.5.2009, 2 males; AO (49°91'N, 14°11'E, 358 chelicera (Fig. 2B). Palps (Fig. 2C): slender, with m a.s.l.), 8.9.2009, 1 deutonymph; MOH (49°91'N, 14°11'E, 415 m a.s.l.), 8.9.2009, 3 tritonymphs; MOH (49°91'N, marked granulation mainly on base of chelal hand 14°09'E, 419 m a.s.l.), 8.9.2009, 2 tritonymphs, 1 proto- and fixed chelal finger, the rest of hand finely nymph; TOH (49°91'N, 14°10'E, 401 m a.s.l.), 8.9.2009, 1 granulate (Fig. 2C); chelal fingers with normal deutonymph; TOH (49°91'N, 14°09'E, 426 m a.s.l.), number of trichobothria (8 on fixed and 4 on mov- 8.9.2009, 5 males, 1 deutonymph, 1 protonymph. able chelal finger), venom apparatus developed Czech Republic, Český Kras, the Šamor-Mramor for- only in movable chelal finger, nodus ramosus est complex (leg. A. Tenčík) – formaldehyde traps situ- close and distal to subterminal trichobothria; ated in: TO (49.89'N, 14.12'E, 462 m a.s.l.), 30.4.–28.5.2008, 1 female; TO (49.90'N, 14.13'E, 466 m a.s.l.), 30.4.– movable chelal finger medially with only 1 acces- 28.5.2008, 3 females; CP (49.89'N, 14.12'E, 452 m a.s.l.), sory tooth and fixed chelal finger medially with 2 28.5.–25.6.2008, 3 females; CP (49.89'N, 14.12'E, 452 m accessory teeth (Fig. 2C). Legs: tarsus IV distally a.s.l.), 25.6.–23.7.2008, 1 female; TO (49.90'N, 14.12'E, 444 with a denticulate seta of highly varying length. m a.s.l.), 23.7.–19.8.2008, 1 female; TO (49.90'N, 14.13'E, Abdominal tergites I–X divided, tergite XI undi- 466 m a.s.l.), 23.7.–19.8.2008, 1 female. vided, tergite XI with a pair of relatively long tac- Slovakia, Malé Karpaty Mts., Cajla (leg. M. Holecová) tile setae (Fig. 2E). Both sexes of species were – sifted from leaf litter in oak-hornbeam forest (48°20'N, 17°16'E, 270 m a.s.l.), 25.5.1999, 1 female (Christophory- measured; body measurements are given in Table ová & Krumpál 2007). 1. Slovakia, Svätý Jur, Šúr Nature Reserve, Panónsky Female (Fig. 2, Table 1): Chaetotaxy of cara- háj (leg. O. Majzlan, Ľ. Vidlička) – Malaise trap situated in pace: carapace with 64–80 setae, 30–37 of them thermophilous old oak forest (48°13'N, 17°12'E, 130 m situated in front of anterior transverse furrow, a.s.l.), 7.6.2008, 1 male (Christophoryová & Krumpál posterior margin with 10–14 setae; 2–4 slitlike lyri- 2010). fissures present in anterior part of carapace, 7–15 All specimens were determined by J. Christophory- ová and K. Krajčovičová using the identification key of lyrifissures behind anterior furrow. Palps (Fig. Christophoryová et al. (2011b). Morphological terminol- 2C): fixed chelal finger with 35–43 and movable ogy follows Beier (1963), Harvey (1992) and Judson chelal finger with 40–51 equally long teeth; fixed (2007), with trichobothrial terminology following Cham- chelal finger laterally with 6–8 accessory teeth and berlin (1931) and taken measurements following Beier movable chelal finger laterally with 5–7 teeth. (1963). Specimens were studied as temporary slide Chaetotaxy of tergites I–X: 10–13 (right tergite half mounts, photographed using a Leica DM1000 compound 4–7: left tergite half 5–6): 10–14 (5–7: 5–8): 10–13 microscope with ICC50 Camera Module (LAS EZ applica- tion, 1.8.0). Measurements were taken from photographs (5–7: 5–6): 10–16 (5–8: 5–8): 13–17 (6–8: 6–9): 12–16 using the AxioVision 40LE application (v. 4.5). The mate- (6–9: 6–8): 13–17 (6–10: 6–8): 13–16 (7–8: 6–9): 12–16 rial is deposited in the zoological collections of the (6–9: 6–8): 11–15 (5–7: 5–8), tergite XI with 7–8 se- Comenius University, Bratislava. tae and with a pair of relatively long tactile setae (Fig. 2E); number of lyrifissures on tergites I–XI: 2– 9: 3–9: 4–9: 4–8: 4–7: 4–8: 5–12: 5–11: 4–7: 2–7: 2. Results Female spermatheca unpaired (Fig. 2D), T- or mushroom-like (apically divided into 2 tubes not Adults (Fig. 2, Table 1): Vestitural setae on body exceeding length of the common stem), female relatively short, dentate and clavate (Fig. 2A); genital operculum anterior with 14–17 setae and 2 carapace (Fig. 2A): granulate and rectangular lyrifissures, operculum posterior with 7–10 setae without cucullus, epistome absent, anterior mar- and 2 lyrifissures. gin of carapace straight, carapace scarcely longer Male (Fig. 2, Table 1): Chaetotaxy of carapace than broad; eyes absent; anterior transverse fur- (Fig. 2A): carapace with 67–87 setae, 32–40 of them 208 J. Christophoryová et al.

Figure 2. Adult of Allochernes peregrinus. A. Carapace with anterior transverse furrow (male, dorsal view). Arrows point to the anterior transverse furrow and one lyrifissure. B. Left chelicera (female, dorsal view). Arrows point to the galea and rallum. C. Left palpal chela with the trichobothrial pattern (female, lateral view). Arrows point to the details of dentition of chelal fingers, granulation of palpal hand and fixed chelal finger and setae. D. Female sper- matheca (ventral view). Arrow points to one lyrifissure. E. Tergit XI (female, dorsal view). Scales: 0.1 (B, D), 0.2 (E) and 0.5 (A, C) mm.

Table 1. Morphometric data for all developmental stages of Allochernes peregrinus (measurements in mm). Abbreviations: M: median, Min: minimum, Max: maximum, n: number of individuals measured, SD: standard deviation, x: arithmetic mean.

Females (n=15) Males (n=15) Tritonymphs (n=10) Deutonymphs (n=6) Protonymphs (n=2) Characteristics Min-Max M/x±SD Min-Max M/x±SD Min-Max M/x±SD Min-Max M/x±SD Min-Max M/x±SD Body, length 1.36-1.97 1.60/1.61±0.14 1.35-1.66 1.47/1.48±0.10 1.15-1.53 1.26/1.29±0.11 1.03-1.24 1.11/1.12±0.08 0.95-0.96 0.96/0.96±0.01 Carapace, length 0.51-0.61 0.56/0.56±0.03 0.50-0.57 0.51/0.52±0.02 0.43-0.49 0.46/0.46±0.02 0.35-0.38 0.37/0.37±0.01 0.32-0.32 0.32/0.32±0.00 Carapace, anterior width 0.31-0.39 0.35/0.35±0.02 0.32-0.35 0.34/0.34±0.01 0.29-0.31 0.30/0.30±0.01 0.23-0.24 0.24/0.24±0.01 0.18-0.18 0.18/0.18±0.00 Carapace, posterior width 0.50-0.61 0.55/0.55±0.03 0.42-0.55 0.45/0.46±0.04 0.36-0.42 0.38/0.38±0.02 0.33-0.36 0.36/0.35±0.01 0.31-0.31 0.31/0.31±0.00 Carapace, length/posterior width ratio 0.98-1.07 1.02/1.03±0.03 1.02-1.21 1.13/1.12±0.05 1.10-1.26 1.20/1.20±0.05 1.00-1.12 1.06/1.05±0.04 1.03-1.03 1.03/1.03±0.00 Chelicera, length 0.19-0.21 0.20/0.20±0.01 0.18-0.21 0.19/0.19±0.01 0.16-0.18 0.17/0.17±0.01 0.13-0.14 0.14/0.14±0.01 0.12-0.12 0.12/0.12±0.00 Chelicera, width 0.08-0.09 0.08/0.08±0.00 0.08-0.09 0.08/0.08±0.00 0.07-0.07 0.07/0.07±0.00 0.06-0.06 0.06/0.06±0.00 0.05-0.05 0.05/0.05±0.00 Chelicera, length/width ratio 2.22-2.63 2.38/2.38±0.11 2.25-2.63 2.38/2.40±0.09 2.29-2.57 2.43/2.40±0.11 2.17-2.33 2.25/2.25±0.09 2.40-2.40 2.40/2.40±0.00 Cheliceral movable finger, length 0.16-0.17 0.16/0.16±0.00 0.15-0.17 0.16/0.16±0.01 0.12-0.14 0.13/0.13±0.01 0.09-0.11 0.10/0.10±0.01 0.08-0.08 0.08/0.08±0.00 Palpal femur, length 0.45-0.52 0.47/0.47±0.02 0.42-0.51 0.43/0.44±0.03 0.28-0.34 0.33/0.32±0.02 0.22-0.24 0.23/0.23±0.01 0.17-0.17 0.17/0.17±0.00 Palpal femur, width 0.14-0.17 0.15/0.15±0.01 0.14-0.16 0.14/0.14±0.01 0.10-0.13 0.12/0.12±0.01 0.09-0.10 0.10/0.10±0.01 0.08-0.08 0.08/0.08±0.00 Palpal femur, length/width ratio 2.76-3.33 3.13/3.12±0.13 2.80-3.40 3.00/3.03±0.14 2.58-2.83 2.75/2.73±0.09 2.30-2.67 2.42/2.43±0.13 2.13-2.13 2.13/2.13±0.00 Palpal patella, length 0.43-0.49 0.45/0.45±0.02 0.39-0.45 0.40/0.41±0.02 0.26-0.31 0.31/0.30±0.02 0.20-0.21 0.21/0.21±0.00 0.17-0.17 0.17/0.17±0.00 Palpal patella, width 0.17-0.20 0.18/0.18±0.01 0.17-0.19 0.18/0.18±0.01 0.13-0.15 0.15/0.15±0.01 0.11-0.11 0.11/0.11±0.00 0.09-0.09 0.09/0.09±0.00 Palpal patella, length/width ratio 2.26-2.67 2.50/2.47±0.12 2.16-2.53 2.35/2.31±0.11 2.00-2.07 2.07/2.05±0.03 1.82-1.91 1.91/1.89±0.04 1.89-1.89 1.89/1.89±0.00 Palpal hand with pedicel, length 0.49-0.57 0.50/0.52±0.03 0.43-0.51 0.45/0.45±0.02 0.33-0.39 0.37/0.37±0.02 0.27-0.29 0.28/0.28±0.01 0.20-0.20 0.20/0.20±0.00 Palpal hand without pedicel, length 0.43-0.50 0.45/0.46±0.02 0.37-0.44 0.38/0.39±0.02 0.30-0.35 0.33/0.33±0.02 0.24-0.26 0.25/0.25±0.01 0.18-0.18 0.18/0.18±0.00 Palpal hand, width 0.23-0.27 0.24/0.24±0.01 0.20-0.24 0.21/0.21±0.01 0.17-0.20 0.19/0.19±0.01 0.14-0.15 0.14/0.14±0.00 0.11-0.11 0.11/0.11±0.00 Palpal hand, length/width ratio 2.04-2.26 2.13/2.14±0.06 2.00-2.20 2.09/2.09±0.07 1.89-2.06 1.95/1.96±0.05 1.71-1.79 1.76/1.75±0.04 1.82-1.82 1.82/1.82±0.00 Palpal (chelal) finger, length 0.38-0.45 0.41/0.41±0.02 0.35-0.38 0.37/0.37±0.01 0.28-0.31 0.30/0.30±0.01 0.22-0.26 0.24/0.24±0.01 0.19-0.20 0.20/0.20±0.01 Palpal chela, length 0.83-0.96 0.87/0.88±0.04 0.76-0.85 0.78/0.78±0.02 0.58-0.67 0.65/0.64±0.03 0.48-0.52 0.49/0.49±0.02 0.38-0.39 0.39/0.39±0.01 Palpal chela, length/palpal hand width 3.41-3.83 3.65/3.63±0.12 3.45-3.95 3.67/3.65±0.15 3.30-3.67 3.38/3.41±0.11 3.43-3.57 3.45/3.47±0.06 3.45-3.55 3.50/3.50±0.07 Leg I femur, length 0.10-0.13 0.11/0.11±0.01 0.09-0.11 0.10/0.10±0.01 0.08-0.09 0.08/0.08±0.01 0.07-0.08 0.07/0.07±0.01 0.06-0.06 0.06/0.06±0.00 Leg I femur, width 0.07-0.09 0.08/0.08±0.01 0.07-0.09 0.08/0.08±0.01 0.06-0.08 0.07/0.07±0.01 0.06-0.06 0.06/0.06±0.00 0.05-0.05 0.05/0.05±0.00 Leg I femur, length/width ratio 1.11-1.57 1.38/1.37±0.13 1.11-1.43 1.25/1.22±0.09 1.13-1.33 1.14/1.19±0.08 1.17-1.33 1.17/1.22±0.09 1.20-1.20 1.20/1.20±0.00 Leg I patella, length 0.19-0.22 0.20/0.20±0.01 0.16-0.19 0.17/0.18±0.01 0.13-0.15 0.14/0.14±0.01 0.11-0.11 0.11/0.11±0.00 0.09-0.09 0.09/0.09±0.00 Leg I patella, width 0.07-0.09 0.08/0.08±0.01 0.07-0.08 0.08/0.08±0.01 0.06-0.07 0.06/0.06±0.00 0.05-0.06 0.06/0.06±0.00 0.04-0.05 0.05/0.05±0.01 Leg I patella, length/width ratio 2.22-2.75 2.50/2.49±0.15 2.13-2.71 2.25/2.32±0.18 2.00-2.33 2.17/2.20±0.13 1.83-2.20 1.83/1.89±0.15 1.80-2.25 2.03/2.03±0.32 Leg I tibia, length 0.19-0.21 0.20/0.20±0.01 0.17-0.20 0.17/0.17±0.01 0.13-0.14 0.14/0.14±0.01 0.10-0.11 0.11/0.11±0.01 0.08-0.08 0.08/0.08±0.00 Leg I tibia, width 0.06-0.07 0.06/0.06±0.00 0.06-0.07 0.06/0.06±0.00 0.05-0.06 0.05/0.05±0.01 0.05-0.05 0.05/0.05±0.00 0.04-0.04 0.04/0.04±0.00 Leg I tibia, length/width ratio 3.00-3.50 3.17/3.19±0.15 2.43-3.33 2.83/2.88±0.19 2.33-2.80 2.60/2.51±0.17 2.00-2.20 2.10/2.10±0.11 2.00-2.00 2.00/2.00±0.00 Leg I tarsus, length 0.20-0.23 0.21/0.21±0.01 0.19-0.22 0.20/0.20±0.01 0.16-0.17 0.17/0.17±0.01 0.13-0.14 0.14/0.14±0.01 0.11-0.12 0.12/0.12±0.01 Leg I tarsus, width 0.05-0.05 0.05/0.05±0.00 0.04-0.06 0.05/0.05±0.00 0.05-0.05 0.05/0.05±0.00 0.04-0.05 0.05/0.05±0.01 0.04-0.04 0.04/0.04±0.00 Leg I tarsus, length/width ratio 4.00-4.60 4.20/4.29±0.24 3.67-5.00 4.00/4.03±0.30 3.20-3.40 3.40/3.32±0.10 2.60-3.50 2.80/2.97±0.42 2.75-3.00 2.88/2.88±0.18 Table 1. (continued)

Females (n=15) Males (n=15) Tritonymphs (n=10) Deutonymphs (n=6) Protonymphs (n=2) Characteristics Min-Max M/x±SD Min-Max M/x±SD Min-Max M/x±SD Min-Max M/x±SD Min-Max M/x±SD Leg IV femur+patella, length 0.37-0.44 0.39/0.39±0.02 0.33-0.37 0.34/0.34±0.01 0.25-0.29 0.28/0.28±0.01 0.20-0.22 0.21/0.21±0.01 0.15-0.16 0.16/0.16±0.01 Leg IV femur+patella, width 0.09-0.11 0.09/0.10±0.01 0.08-0.10 0.09/0.09±0.01 0.07-0.08 0.08/0.08±0.01 0.06-0.07 0.06/0.06±0.00 0.05-0.05 0.05/0.05±0.00 Leg IV femur+patella, length/width ratio 3.73-4.56 4.11/4.11±0.23 3.40-4.25 3.89/3.91±0.27 3.50-3.71 3.54/3.57±0.08 3.14-3.50 3.33/3.36±0.13 3.00-3.20 3.10/3.10±0.14 Leg IV tibia, length 0.25-0.31 0.27/0.27±0.02 0.22-0.27 0.24/0.24±0.01 0.16-0.19 0.19/0.18±0.01 0.13-0.15 0.14/0.14±0.01 0.09-0.10 0.10/0.10±0.01 Leg IV tibia, width 0.06-0.07 0.07/0.07±0.00 0.06-0.07 0.06/0.06±0.01 0.05-0.06 0.06/0.06±0.00 0.05-0.06 0.06/0.06±0.01 0.05-0.05 0.05/0.05±0.00 Leg IV tibia, length/width ratio 3.57-4.43 3.86/3.96±0.26 3.29-4.17 3.83/3.75±0.28 3.00-3.20 3.17/3.12±0.08 2.33-2.80 2.50/2.51±0.18 1.80-2.00 1.90/1.90±0.14 Leg IV tarsus, length 0.25-0.28 0.25/0.26±0.01 0.23-0.24 0.24/0.24±0.01 0.17-0.20 0.20/0.19±0.01 0.15-0.16 0.15/0.15±0.01 0.12-0.12 0.12/0.12±0.00 Leg IV tarsus, width 0.06-0.06 0.06/0.06±0.00 0.05-0.06 0.05/0.05±0.00 0.05-0.06 0.06/0.06±0.00 0.04-0.05 0.04/0.04±0.01 0.04-0.04 0.04/0.04±0.00 Leg IV tarsus, length/width ratio 4.17-4.67 4.17/4.28±0.16 4.00-4.80 4.60/4.49±0.32 3.17-3.40 3.33/3.27±0.09 3.20-3.75 3.75/3.57±0.28 3.00-3.00 3.00/3.00±0.00

Figure 3. Tritonymph of Allochernes peregrinus. A. Detail of cheliceral galea (dorsal view). Figure 4. Deutonymph of Allochernes peregrinus. A. Detail of cheliceral galea (dorsal view). B. Left palpal chela with the trichobothrial pattern (lateral view). C. Tergit XI (dorsal B. Left palpal chela with the trichobothrial pattern (lateral view). C. Tergit XI (dorsal view). Scales: 0.01 (A), 0.1 (C) and 0.2 (B) mm. view). Scales: 0.01 (A), 0.1 (C) and 0.2 (B) mm.

Description of developmental stages of Allochernes peregrinus 211

Figure 5. Protonymph of Allochernes peregrinus. A. Detail of cheliceral galea (dorsal view). B. Left pal- pal chela with the trichobothrial pattern (lateral view). C. Tergit XI (dorsal view). Scales: 0.01 (A), 0.1 (C) and 0.2 (B) mm.

situated in front of anterior transverse furrow, Tritonymphs (Fig. 3, Table 1): Chaetotaxy of posterior margin with 10–12 setae; 2 slitlike lyrifis- carapace: carapace with 56–64 setae, 29–33 of them sures present in anterior part of carapace, 6–12 situated in front of anterior transverse furrow, lyrifissures behind anterior furrow (Fig. 2A). posterior margin with 10–12 setae; 2 slitlike lyrifis- Palps: fixed chelal finger with 33–42 and movable sures present in anterior part of carapace, 3–7 lyri- chelal finger with 39–44 equally long teeth; fixed fissures behind anterior furrow. Chelicera: 5 setae chelal finger laterally with 5–9 accessory teeth and on cheliceral hand, one on cheliceral movable fin- movable chelal finger laterally with 5–7 teeth. ger; movable cheliceral finger with slender well- Chaetotaxy of tergites I–X: 10–13 (right tergite half developed galea, main stalk with approximately 5–7: left tergite half 5–6): 10–12 (5–6: 4–7): 9–11 (5– 4–5 short terminal rami (Fig. 3A). Palps (Fig. 3B): 7 6: 4–5): 10–14 (5–7: 5–7): 11–14 (5–8: 6–7): 12–16 (6– trichobothria on fixed chelal finger and 3 on mov- 8: 6–8): 11–15 (6–8: 5–8): 12–15 (6–7: 6–8): 12–14 (6– able chelal finger; fixed chelal finger with 27–30 7: 6–7): 9–14 (4–7: 4–8), tergite XI with 6–8 setae and movable chelal finger with 31–35 equally long and with a pair of relatively long tactile setae; teeth; fixed chelal finger laterally with 4–6 acces- number of lyrifissures on tergites I–XI: 2–5: 2–6: 2– sory teeth and movable chelal finger laterally with 7: 3–7: 3–7: 3–6: 3–7: 3–5: 2–5: 2–4: 2. Male genital 4–5 teeth (Fig. 3B). Chaetotaxy of tergites I–X: 10– operculum anterior with 15–19 setae and 2 lyrifis- 11 (right tergite half 5–6: left tergite half 5–6): 9–11 sures, operculum posterior with 8–9 setae. (4–6: 4–6): 9–10 (4–5: 4–5): 10–12 (4–6: 5–6): 10–14 (5–8: 5–6): 10–14 (5–7: 5–7): 10–13 (5–7: 5–7): 10–15 Nymphs (Fig. 3–5, Table 1): In most aspects the (5–8: 5–7): 8–15 (3–8: 5–7): 8–12 (3–6: 4–6), tergite morphology of tritonymphs, deutonymphs and XI with 6 setae and with a pair of relatively long protonymphs is similar to that of adults (e.g. mor- tactile setae (Fig. 3C); number of lyrifissures on phology of vestitural setae on body, granulation of tergites I–XI: 2–5: 2–4: 3–5: 2–4: 2–4: 3–5: 2–4: 2–4: carapace, absence of epistome and eyes, morphol- 2–4: 2–3: 2. ogy of chelicerae, morphology of palps, presence Deutonymphs (Fig. 4, Table 1): Chaetotaxy of of venom apparatus in movable chelal finger, carapace: carapace with 45–53 setae, 24–28 of them presence of a pair of relatively long tactile setae on situated in front of anterior transverse furrow, tergite XI). Body measurements are given in Table posterior margin with 9–10 setae; 2 slitlike lyrifis- 1. sures present in anterior part of carapace, 1–3 lyri- 212 J. Christophoryová et al. fissures behind anterior furrow. Chelicera: 5 setae Gabbutt 1970). on cheliceral hand, one on cheliceral movable fin- The first description of Allochernes peregrinus ger; movable cheliceral finger with slender well- was based on the following characters of one fe- developed galea, main stalk with approximately 4 male specimen – chaetotaxy of posterior carapace short terminal rami (Fig. 4A). Palps (Fig. 4B): 6 margin and pedal tarsus IV, lateral teeth of chelal trichobothria on fixed chelal finger and 2 on mov- fingers, cheliceral galea, rallum and palpal meas- able chelal finger; fixed chelal finger with 25–29 urements (Lohmander 1939). Lohmander (1939) and movable chelal finger with 27–32 equally long drew figures of palpal chela, leg IV and cheliceral teeth. Chaetotaxy of tergites I–X: 6–10 (right tergite galea. Later Mahnert (1983) gave some data of one half 3–5: left tergite half 3–5): 7–9 (3–5: 3–5): 6–9 Hungarian female specimen – measurement of (3–5: 3–5): 7–9 (3–4: 4–5): 8–10 (4–5: 4–5): 8–10 (4–5: palpal chela, leg I and IV, number of teeth of che- 4–5): 8–10 (4–5: 4–5): 8–10 (4–5: 3–5): 8–9 (4–5: 3–5): lal fingers (including medial and lateral accessory 6–8 (3–4: 3–4), tergite XI with 6 setae and with a teeth) and drew palpal chela with the pair of relatively long tactile setae (Fig. 4C); num- trichobothrial pattern and female spermatheca. ber of lyrifissures on tergites I–XI: 2–4: 2–4: 2–4: 2– Further data are partly available in the theses of 4: 2–3: 2–4: 2–4: 2–3: 2–3: 2: 2. Ducháč (2002) and DeVore-Scribante (1999) Protonymphs (Fig. 5, Table 1): Chaetotaxy of mainly concerning females, but also chaetotaxy of carapace: carapace with 27 setae, 17–18 of them male genital operculum anterior and chelicera and situated in front of anterior transverse furrow, palps of males and tritonymphs. DeVore-Scribante posterior margin with 6 setae; 2 slitlike lyrifissures (1999) added also figures of chelicera and inaccu- present in anterior part of carapace, 2–3 lyrifis- rate figure of carapace. However these data and sures behind anterior furrow. information were not published. Chelicera: 4 setae on cheliceral hand, cheliceral The main taxonomic characters of the studied movable finger without seta; movable cheliceral females correspond to published descriptions finger with slender well-developed galea, main from Sweden (Lohmander 1939) and Hungary stalk with approximately 3 short terminal rami (Mahnert 1983), despite registering greater vari- (Fig. 5A). Palps (Fig. 5B): 3 trichobothria on fixed ability in morphometric characteristics of the chelal finger and 1 on movable chelal finger; fixed body, carapace, palps, legs I and leg IV. Compared chelal finger with 21–23 and movable chelal finger to the known description, we recorded smaller with 24–25 equally long teeth. Chaetotaxy of all and bigger specimens as well. We registered a tergites from I to X: 6–6 (right tergite half 3–3: left higher variability also in the number of marginal tergite half 3–3), tergite XI with 2 setae and with a teeth and lateral accessory teeth. For the first time pair of relatively long tactile setae (Fig. 5C); num- we present female chaetotaxy and number of lyri- ber of lyrifissures on tergites I–XI: 2–3: 2–3: 2–4: 2– fissures of carapace, genital operculum anterior, 4: 2–3: 2–3: 2–3: 2–2: 2–3: 2–3: 2. genital operculum posterior, all tergites and cheli- ceral measurements. Five Allochernes species and subspecies are Discussion currently known from neighbouring countries (Table 2): A. bulgaricus Hadži, 1939, A. mahnerti The taxonomy and identification of pseudoscor- Georgescu & Capuse, 1996, A. powelli (Kew, 1916), pion species often use morphometric characters, A. wideri phaleratus (Simon, 1879) and A. wideri but are frequently based on limited material, wideri (C.L. Koch, 1843). The species A. peregrinus mostly on a few type specimens (especially in differs from the others by presence of a pair of Beier 1963). Under these circumstances, only a relatively long tactile setae on tergite XI (Table 2). poor reflection of intraspecific variability is avail- From A. mahnerti (Georgescu & Capuse 1996), it able (Christophoryová et al. 2011b). Detailed mor- differs by longer palpal femur, longer chelal fin- phological or morphometric analysis of common ger, higher number of teeth on chelal fingers, species are mostly rare and typical mainly for re- higher number of lateral accessory teeth and in cent descriptions of new taxa. The published data chaetotaxy of female genital operculum (Table 2). about descriptions of all developmental stages of From other species and subspecies (Beier 1932, European pseudoscorpion species are mainly lim- 1963, Hadži 1939, Helversen 1966, Legg & Jones ited to Chthoniidae, Neobisiidae and Cheliferidae 1988, Christophoryová et al. 2011a), A. peregrinus (e.g. Gabbutt & Vachon 1963, 1965, 1967, 1968, differs also in number of lateral accessory teeth

Description of developmental stages of Allochernes peregrinus 213

Table 2. The comparison of adults of Allochernes species and subspecies (measurements in mm). Abbreviations: +: presence, –: absence, ?: unknown data.

Characteristics/ taxon peregrinus bulgaricus mahnerti powelli phaleratus wideri wideri wideri ...... A A A A A A

Presence of tactile setae on tergit XI + – – – – – Length of palpal femur 0.42-0.52 0.52 0.34-0.37 0.58-0.85 0.70 0.63-0.77 Length of chelal finger 0.35-0.45 0.37 0.25-0.27 0.44-0.55 0.54 0.49-0.59 Number of teeth on fixed chelal finger 33-43 32 29 35-40 42 41 Number of teeth on movable chelal finger 39-51 37 32 31-41 39 43-44 Number of lateral accessory teeth on fixed chelal finger 5-9 4 3 3-4 4 2-4 Number of lateral accessory teeth on movable chelal finger 5-7 1 3 4-5 3 2-4 Number of setae on female genital operculum anterior 14-17 ? 22 28-34 ? 20-22 Number of setae on female genital operculum posterior 7-10 ? 11 8 ? 9-12 Number of setae on male genital operculum anterior 15-19 22+2 19+6 25 ? 35-37 Number of setae on male genital operculum posterior 8-9 17 10 17-22 ? 20-21

and chaetotaxy of female or male genital opercu- tritonymphs and deutonymphs in May and lum (Table 2). September and protonymphs in September. It was The published records of A. peregrinus came remarkable that only females were present in the from specimens found (mainly individually) in the formaldehyde traps. Other developmental stages deeper soil layers under sunken stones (Rafalski were found after collecting of the soil samples. 1967, Krumpálová & Krumpál 1993) or specimens Until now, only isolated findings of A. peregrinus were collected from soil samples and sifted from were recorded in the Czech Republic (Ducháč leaf and coniferous litter (Muchmore 1972, 1990, 1995, 1999, 2002). Accordingly, the species was Drogla 1983, Ducháč 1995, Drogla & Lippold 2004, listed in the Red Data Book of the Czech Republic Christophoryová & Krumpál 2007). In the Czech as vulnerable (Šťáhlavský & Ducháč 2005). Our Republic A. peregrinus was swept from vegetation research refers that the Český kras Protected (Ducháč 2002), and in Slovakia it was collected Landscape Area presents an appropriate locality using a Malaise trap situated in an oak forest with habitats and conditions suitable for (Christophoryová & Krumpál 2010). Several occurrence of stable and relatively numerous specimens were phoretic and were attached to legs populations of A. peregrinus. of a fly Delia floralis (Lohmander 1939) or harvestmen Opilio canestrinii (Thorell, 1876), Leiobunum rupestre (Herbst, 1799) and L. rotundum (Latreille, 1798) (Drogla & Lippold 2004). Adults Acknowledgements. We are grateful to our colleagues occurred from May to October (Lohmander 1939, Aleš Tenčík, Milada Holecová, Ľubomír Vidlička and Oto Muchmore 1972, Drogla 1983, Mahnert 1983, Majzlan for collecting pseudoscorpion material, Alica Krumpálová & Krumpál 1993, Drogla & Christophoryová for technical assistance with figures and Lippold 2004, Christophoryová & Krumpál the reviewers Dr. Gardini, Dr. Zaragoza and Dr. 2010). The finding of A. peregrinus during our Štáhlavský for their valuable comments and critic. The study was supported by the projects VaV SP/2d3/139/07 research in the Český kras Protected Landscape (Limits of biodiversity protection in fragmented Area corresponds to the known ecological landscape) of the Ministry of Environment of the Czech demands of this species. The specimens occurred Republic and the project of VEGA 1/0176/09. in oak and oak-hornbeam forests and plantations of deciduous and coniferous trees from 234 to 466 m a.s.l. Females were collected from April to August, males in May, June and September, 214 J. Christophoryová et al.

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